September 24, 2009

Modern Scandinavians descended (maybe) from Neolithic TRB but not Mesolithic Pitted Ware ancestors

Coming shortly after Bramanti et al. (2009) which discovered a discontinuity between Neolithic farmers from Central and Eastern Europe and the pre-existing hunter-gatherers, a new study examines ancient DNA from northern European populations, extending the picture of discontinuity all the way to Scandinavia. This is one more nail in the coffin of the cultural diffusion hypothesis, and in favor for a demic diffusion of agriculture all the way to the northernmost reaches of Europe.

More on this after I read the full paper.

UPDATE (Sep 25):

From the paper:

Although the hunter-gatherers of Denmark and southern Sweden adopted pottery early on, the Neolithization first took real shape with the appearance of the Funnel Beaker Cultural complex (FBC, also known as the Trichterbecher Kultur [TRB]) some 6,000 years BP (the oldest evidence possible dating back some 6,200 years BP [9]). Atthis time domestic cattle and sheep, cereal cultivation, and the characteristic TRB pottery were introduced into most of Denmark and southern parts of Sweden [6]. Nevertheless,the Neolithization process was slow in Scandinavia, and large are as remained populated by hunter-gatherer groups until the end of the 5th millennium BP.

One of these last hunter-gatherer complexes was the Pitted Ware culture (PWC), which can be identified by its single-inhumation graves distributed over the coastal areas of Sweden and the Baltic Sea islands that lie closest to the Swedish coast. Intriguingly, the PWC first appears in the archaeological record of Scandinavia after the arrival of the TRB (some 5,300 yearsBP) and existed in parallel with farmers for more than a millennium before vanishing about 4,000 years BP (Figure 1).

The authors sampled 3 TRB individuals from "one passage tomb, Gokhem, dated to 5,500–4,500 years BP" which were found to belong to haplogroups H, J, and T, and 19 PWC individuals "from three different sites on the Baltic island of Gotland dated to 4,800–4,000 years BP" which were found to belong to haplogroups J, T, V (one each), "Other" (two), U5 and U5a (three each), and U4/H1b (eight samples).

From the paper:

Given our results, it remains possible that the PWC represent remnants of a larger northern European Mesolithic hunter gather complex. However, it appears unlikely that population continuity exists between the PWC and contemporary Scandinavians or Saami. Thus, our findings are in agreement with archaeological theories suggesting Neolithic or post-Neolithic population introgression or replacement in Scandinavia.To what extent this holds true for other parts of Europe requires further direct testing, although morphological [24, 25], ancient [26], and modern [4, 5] genetic data suggest that this is probably the case.

The results indicate that the PWC was dominated by haplogroup U (about three quarters of the mtDNA gene pool). The inability to resolve between U4 and H1b is due to the portion of the mtDNA sampled. Given (i) the absence of other H subgroups in the large sample, (ii) the higher frequency of U4 in modern populations, (iii) the presence of U4 but not H1b in other pre-farming populations of Europe (after Bramanti et al.), (iv) the absence of U4 in Neolithic populations, (v) the higher coalescence age of U4 compared to H1b, suggesting a deeper ancestry, I am inclined to think that most, if not all of the U4/H1b is actually just U4.

UPDATE II:

Fst between the PWC and modern populations ranged between 0.036 (Latvians) and Saami (0.25). For Swedes and Norwegians they were 0.051 and 0.061. A few conclusions can be drawn from this:

The notion of Saami as unmixed descendants of pre-farming Europeans is debunked.

Latvians and other populations of the eastern Baltic are the closest (although by no means very close) to the PWC.

Swedes and Norwegians are somewhat closer to the pre-farming inhabitants than is the case for Central Europe where Fst=0.086 was estimated by Bramanti et al. (2009)

Traditional physical anthropology held that there were three main elements in northern Europe, which have been given different names, but can be summarized as follows:

Narrow- and high-faced populations, a new element in the region, similar to that of Central Europe

Latvia's most ancient inhabitants tended to be large in size, with large skulls, a distinctly oblong head shape, a broad, high face and a distinctly protruding nose (Denisova 1975). Looking at this data in the context of synchronous populations elsewhere in Europe, we can find specific geographic differentials. This is especially true of the facial width of residents, a factor which has great weight in the specification of race (Denisova 1978). Differences in facial width in Europe became particularly distinctive at the beginning of the Atlantic period, when farming was begun in Europe. At this time, facial width distinctly separated morphological forms in Northern Europe from those in the Mediterranean region -- two distinct geographic regions. Massive, broad-faced morphological forms dominated in northern and northeastern Europe, while gracile, narrow-faced forms are found most often in Middle Europe and the continent's southeastern reaches. During the Atlantic period, narrow-faced populations gradually moved in the northerly and northeasterly direction. They reached the Baltic region only during the Bronze Age. For this reason, during the Mesolithic and Neolithic period, people in the Baltic region (and surrounding regions) had broad faces, a fact which affirms their links to the late Paleolithic populations of Europe.

Modern Scandinavians are more (1) than (2), while modern Balts are more (2) than (1). The mtDNA picture seems fairly consistent with a greater persistence of Proto-Europoid elements among the Balts.

Today's Scandinavians are not descended from the people who came to Scandinavia at the conclusion of the last ice age but, apparently, from a population that arrived later, concurrently with the introduction of agriculture. This is one conclusion of a new study straddling the borderline between genetics and archaeology, which involved Swedish researchers and which has now been published in the journal Current Biology.

"The hunter-gatherers who inhabited Scandinavia more than 4,000 years ago had a different gene pool than ours," explains Anders Götherström of the Department of Evolutionary Biology at Uppsala University, who headed the project together with Eske Willerslev of the Centre for GeoGenetics at the University of Copenhagen.

The study, a collaboration among research groups in Sweden, Denmark and the UK, involved using DNA from Stone Age remains to investigate whether the practices of cultivating crops and keeping livestock were spread by immigrants or represented innovations on the part of hunter-gatherers.

"Obtaining reliable results from DNA from such ancient human remains involves very complicated work," says Helena Malmström of the Department of Evolutionary Biology at Uppsala University.

She carried out the initial DNA sequencings of Stone Age material three years ago. Significant time was then required for researchers to confirm that the material really was thousands of years old.

"This is a classic issue within archaeology," says Petra Molnar at the Osteoarchaeological Research Laboratory at Stockholm University. "Our findings show that today's Scandinavians are not the direct descendants of the hunter-gatherers who lived in the region during the Stone Age. This entails the conclusion that some form of migration to Scandinavia took place, probably at the onset of the agricultural Stone Age. The extent of this migration is as of yet impossible to determine."

The driving force behind the transition from a foraging to a farming lifestyle in prehistoric Europe (Neolithization) has been debated for more than a century [1], [2] and [3]. Of particular interest is whether population replacement or cultural exchange was responsible [3], [4] and [5]. Scandinavia holds a unique place in this debate, for it maintained one of the last major hunter-gatherer complexes in Neolithic Europe, the Pitted Ware culture [6]. Intriguingly, these late hunter-gatherers existed in parallel to early farmers for more than a millennium before they vanished some 4,000 years ago [7] and [8]. The prolonged coexistence of the two cultures in Scandinavia has been cited as an argument against population replacement between the Mesolithic and the present [7] and [8]. Through analysis of DNA extracted from ancient Scandinavian human remains, we show that people of the Pitted Ware culture were not the direct ancestors of modern Scandinavians (including the Saami people of northern Scandinavia) but are more closely related to contemporary populations of the eastern Baltic region. Our findings support hypotheses arising from archaeological analyses that propose a Neolithic or post-Neolithic population replacement in Scandinavia [7]. Furthermore, our data are consistent with the view that the eastern Baltic represents a genetic refugia for some of the European hunter-gatherer populations.

47 comments:

The study's title doesn't say Scandinavians are descended from Neolithic farmers, like you put it. The study said the ancient mtdna is closer to modern Baltic people. I didn't even know there was a difference in the mtdna of Baltic and Scandinavian people, this will be curious...

By the way, these remains are from just 4000 years ago. When Maju pointed out that paleolithic Iberians had lots of mtdna H you said they could be Neolithic people because they were within the Neolithic timeframe (Maju corrected you that the Neolithic started later in southwest Europe and the remains were from before that time period in Iberia). If these Scandinavian results included lots of H, I presume you would have immediately rejected their status as paleolithic people and claimed that by 4000 years ago everybody in Europe must have been heavily intermixed with the Neolithic farmers, anyhow.

Talk about ignoring the huge Mammoth (hunter) in the room. This paper is a triumph of bias over good sense.

There are only 3 Funnel beaker individuals, Haplogroup H, J and T identified. And presumably 16 Pitted Ware People. The Pitted Ware represent the Hunter Gathers and the Funnel Beaker people the Farmers.

Modern Swedish populations (also quoted in paper) visibly correlate much more significantly with the Pitted Ware population (Hunter Gatherers).

Haplogroup H is a known Hunter Gatherer PROVING that hunter gathers embraced agriculture (the reverse of the papers conclusion). H subclades left behind in the east (H6, H8) are not found.

The other two Haplogroups (J and T) are also found in the parallel Pitted Ware population and at levels similar to the modern Swedish population. Indicating that the Hunter Gathers prevailed (reverse conclusion of the paper).

Two Hunter -Gather (U) strains were absent from the farming sample (SAMPLE OF 3 PEOPLE)!

The samples are from an area that is not part of the known Saami territory so can draw NO conclusions about the Saami.

If these Scandinavian results included lots of H, I presume you would have immediately rejected their status as paleolithic people and claimed that by 4000 years ago everybody in Europe must have been heavily intermixed with the Neolithic farmers, anyhow.

No need for counterfactuals. The Pitted Ware people did not include lots of H, while 1 in 3 TRB samples did. The results are entirely consistent with the picture from Central Europe. In particular, the Pitted Ware sample is U-dominated and shows no evidence of the wide assortment of haplogroups present in modern Europeans. The implication is clear that the Pitted Ware hunter-gatherers are not the main ancestors of modern Scandinavians.

As for the idea of an eastern Baltic refugium, this may be interesting to consider, but we now know that U4 and U5 were spread all over Central and Eastern Europe before the advent of the Neolithic, so there is nothing specifically "eastern Baltic" about them.

Haplogroup H is a known Hunter Gatherer

Unfounded premise. H has not been positively documented in ~40 Mesolithic/Paleolithic individuals from the combined sample of this paper and Bramanti et al. (2009). On the contrary, it has been documented in the TRB, LBK, Corded Ware, and Iron Age Lichtenstein.

The evidence is stacked against the idea that H is a "known hunter-gatherer" and in favor of the idea that it was brought by farmers.

Unless Paleolithic Cromagnon (circa 24,000 BCE) had a secret vegetable plot Haplogroup H is a Hunter-Gatherer type. To my knowledge its very origin age (which I think is an underestimate) makes it a Hunter-Gather type. No sign of agriculture 30,000 years ago.

Personally I thing the Bramanti paper is interesting. But the raw data warrants looking at. Most of the Mesolithic samples that came out were very northern geographically. The bulk of the German (Ostorf) samples are Funnel Beaker Culture (farming, 3,000 BC) but classed as Mesolithic presumably because the samples are rich in Haplogroup U and other clearly Hunter Gatherer types.(we dont want to mess up the story do we).

The chief marker of the migration of the farmers from the east, Haplogroup J is only found in 2 samples. Only one in the Neolithic group. The other in the "mesolithic" funnel beaker people.

This paper shows almost no Eastern types associated with early farming in Europe. Very, very little migration from the east.

What it may show is conversion of southern-Hunter gatherers to farming followed by a huge south to north population explosion as the climate improved.

I don't see how the introduction of new mates would mean that the male gene pool was overrun. It is quite peculiar that today there is no male ancestor living --- anywhere, that pre-dates the M253 mutation. Perhaps hunter-gatherer Scandinavia was hit by at least two waves, or three including far northern Russian immigrants. One being from the Baltic region, and the other from the Atlantic. If we break down these 3 major lineages, we have I1(and I2b lines), R1b, and R1a. That would be no different from what is found in Scandinavia today. It doesn't seem like the population was "replaced" at all.

Dienekes argues like a lawyer, not like a scientist. He wants to prove his case, for some reason that isn't scientific, and he points out only the things that are in favor of it, "forgetting" the ones that contradict it, even claiming the ancient Iberian mtdna study is bogus (right, the authors made it up, it's a total fabrication, they're just lying). He constantly refers to things that are very questionable as if they were an established fact, like when he talks about the Aryan Invasion Theory as if it were the Aryan Invasion Fact.

The study of Neolithic mtdna by Haak found 6 out of 24 N1a, spread out over different sites and with 5 different haplotypes. So N1a was supposedly a huge part of the farmers, in fact, the single most important haplogroup. Today N1a is 0,25% of Europe's mtdna. On the other hand, the hunter-gatherers supposedly were mainly U4 and U5, which wasn't found at all in the farmers. In north Europeans today, U4/U5 is 10% to 15% of their mtdna. In fact, Haak's original conclusion was that modern Europeans aren't descended from the Neolithic farmers. The only way to somehow reconcile these bizarre results is to pull the evolution trump card: N1a must be less advantaged than other mtdna haplogroups. I think the notion that different haplogroups have different evolutionary advantages is very far fetched; not impossible, but very far fetched. It's also funny how N1a would have been the only haplogroup that had this evolutionary disadvantage, the others were all exactly equally matched in the evolutionary game. Because the rest of the Haak haplogroups are mainly in line with the modern mtdna of Caucasians. The same thing happens with the ancient mtdna studies of Iberians and Moroccans: their results are similar to modern mtdna. So evolution didn't affect all those haplogroups, they remained unchanged in their relative ratios for thousands of years, only N1a seemed to have an evolutionary disadvantage, and a huge one which took it from being the most important haplogroup to virtually extinct. It's also curious that N1a wasn't found in the ancient Iberian or Moroccan mtdna, given that it was the single most important haplogroup in Haak's study.

Here's a curious consequence to the farmer expansion theory. Why did the farmers that reached north Europe evolve blond hair and blue eyes? They came from the Middle East or Anatolia (remember that Dienekes' himself ruled out their origin in south Europe because it was too unlikely that people living in such close proximity would have such differing sets of haplogroups without mixing), so they were all black haired and brown eyed. And then they magically transformed themselves into mostly blond haired and blue eyed people over just a few thousand years. We know that by Roman times they were already described as being blond haired. So they evolved blond hair and blue eyes in a few thousand years. That's pretty funny. I wonder what are the evolutionary advantages of farming with blond hair and blue eyes in north Europe? They better be real solid advantages, too, because the change was pretty huge, and in just a couple thousand years. And it didn't happen in the south, how funny. This evolutionary advantage only affects farming in the north...

Here's a map of the modern distribution of H, which fits absolutely perfectly with ancient geneflow and genetic continuity. Notice how the percentages gradually decline as one travels away from Europe in all directions. Did all this happen between 5000 and 2000 years ago?? Seems extremely unlikely. Notice how perfectly the percentages gradually decrease starting from the point of geneflow between Europe and North Africa, the Strait of Gibraltar. Notice how they gradually decline from Europe through Anatolia, then Egypt, Nubians, and finally Ethiopia (Sudan had 5% H/HV). Everything about this map points to a very ancient presence of H in Europe, with geneflow resulting in gradually decreasing clines across the 3 routes out of Europe (Gibraltar, Anatolia/Caucasus, the Ural region). How does one explain this map with evolution? Or with an origin of H in the Middle East?

Here's a curious consequence to the farmer expansion theory. Why did the farmers that reached north Europe evolve blond hair and blue eyes? They came from the Middle East or Anatolia (remember that Dienekes' himself ruled out their origin in south Europe because it was too unlikely that people living in such close proximity would have such differing sets of haplogroups without mixing), so they were all black haired and brown eyed.

Well, you may think that it is pretty funny, but the evidence, such as it is, does suggest that light pigmentation has evolved relatively recently in Caucasoids, within a roughly Neolithic time frame.

Also, there is no reason to think that people from Anatolia and the Middle East were all black haired and brown eyed at the time when they entered Europe.

The study of Neolithic mtdna by Haak found 6 out of 24 N1a, spread out over different sites and with 5 different haplotypes. So N1a was supposedly a huge part of the farmers, in fact, the single most important haplogroup. Today N1a is 0,25% of Europe's mtdna. On the other hand, the hunter-gatherers supposedly were mainly U4 and U5, which wasn't found at all in the farmers. In north Europeans today, U4/U5 is 10% to 15% of their mtdna.

Bramanti et al. determine that modern populations are indeed closer (fst=0.058) to the LBK than they are to the hunter-gatherers (0.0858).

Modern Central Europeans' descent from the LBK requires only negative selection against N1a.

Their descent from pre-farming populations can be positively excluded, as pre-farming populations lacked (or had in low frequency) most of the haplogroups found in modern Europeans. Such a descent would require that

(i) the non-detected haplogroups did in fact exist at low frequencies(ii) that they increased in frequency over time to their present-day levels(iii) that correspondingly the U- types decreased in frequency over time.

Quite clearly descent from the LBK is a hell of a lot more parsimonious than descent from the hunter-gatherers.

"At any rate, the finding of the Cambridge Reference Sequence in Paglicci 23 shows that one of today's mtDNA variants has been present in Europe for at least 28,000 years". The Cambridge Reference sequence is Haplogroup H (H2a2).

Caramelli says (2003):

"The D-loop sequence of Paglicci-25 shows no substitutions withrespect to the Cambridge reference sequence (28)."

What it may show is conversion of southern-Hunter gatherers to farming followed by a huge south to north population explosion as the climate improved.

Of course. But not so much the climate. First, it took a long time to have sereals and animals adapted to the colder North with its unsuitable wet summers (that destroy unsuitable crops), and before people knew how to construct proper buildings to protect the animals during the longer winters, and how to properly store feed for them. In addition, there are vast stretches of very poor, sandy soils in the northern plains.

I took about 500 years for agriculture to become successful in the middle Danube region north/northwest of the Balkans. After suitable grains and animals were present, and the required technologies were established, agriculture indeed spread very quickly over much of central Europe, and north along the clay/loess soil river valleys (Elbe/Saale, Weser, Rhein) all the way to nowadays Holland, Belgium, and northeastern France. But the vast regions of sandy soils between Holland and Poland were not initially settled, and likely acted as a barrier for agriculturalists moving into Denmark and Sweden - until population pressure forced people to adopt and adapt to even the poorer soils.

Farther east the soils are so heavy and waterlogged that agriculture also could only progress a little later, perhaps after better drainage systems had been developed.

"At any rate, the finding of the Cambridge Reference Sequence in Paglicci 23 shows that one of today's mtDNA variants has been present in Europe for at least 28,000 years". The Cambridge Reference sequence is Haplogroup H (H2a2).

Your inference is incorrect. Most of rCRS mtDNA may be in haplogroup H today but the rCRS is not exclusive to haplogroup H (or even more specifically H2a2).

If we look at a rCRS of an individual living today, we can infer with great probability that he belongs to haplogroup H. But, this is due to the fact that most rCRS today is in haplogroup H. In the absence of information about the distribution of mtDNA haplogroups 28,000 years ago, that inference is unfounded.

To make the case clear, consider the modern distribution of the rCRS (xls). The rCRS occurs in diverse haplogroup backgrounds: U*, HV*, H, and V. Its frequency in these backgrounds is: 0.1%, 5.4%, 6.4%, 3.8%.

Today's population has 15.5 times more H than HV*. Thus, an rCRS is 6.4*15.5/5.4=18 times more likely to belong to H than to HV*.

The same cannot be said for the population living 28,000 years ago. For one thing, the paucity of mtDNA haplogroup H in the combined available pre-farming sample is much lower than in the modern population. For another, the coalescence time for H is much later than 28,000 years, making it likely that the H clade was not even in existence 28,000 years ago.

But, even if one were to reject both these arguments, the fact remains that in the absence of information about the haplogroup distribution 28,000 years ago, the inference that it was similar to that of today is false, and has been decisively proven false by the two recent studies which showed discontinuity between a few thousand years and today, let alone 28,000 years and today.

"The notion of Saami as unmixed descendants of pre-farming Europeans is debunked."

I have read the paper, that is not what the paper says. The Pitted Ware culture do not overlap with the current saami settlement and overlap just some of the earlier known settlement, we also only know the genetics of the saami from the northern shield of Fennoscandia and not anything of the ancient saami groups further south that dissappeared as the result of scandinavian and finnic expansion. There is however some itneresting finds in southern saami language that supports baltic language contacts not seen in other saami languages that may suggest southern saami language arrived from the south not from the north.

I have never accepted the Paleolithic/Neolithic paradigm to the peopling of Europe. It is a more of a religion than any scientific facts, assumptions based on shards, certain few remains and the finder's belief system.

The tests on ancient human dna from European remains has shown that no modern Europeans whether light or dark, tall or short, fat or thin, Southern or Northern can be linked by dna to either the Mesolithic pre farming populations or the Neolithic era farming populations that lived in Europe. Scandinavia whatever the ethnicity there, is exactly the same as the rest of Europe, unconnected to the Mesolithic hunter/gatherers or Neolithic farmers. The answer to who the ancestors of modern Europeans are has to do with the spreading of Indo-European languages into Europe and the Y chromosome haplogroups R1b and R1a. Mitochrondrial haplogroups H1, H3 and V have been shown to be European in origin. Haplogroups belonging to U while ancient cannot be proven to have originated in Europe but may have been carried into Europe along with the H*.

If you are going to argue based on results you have to do so accurately. For instance, the bones found in Italy are not Cro Magnon. Cro Magnons existed in France. Those ancient Italians are just one of the many AMH who lived in Europe. The Italian remains are best referred to as Cro Magnoid due to their very superficial resemblance, mainly thicker bones. Ancient Europeans went from the elongated, tropical African skeletal structure found in the Cro Magnons to a short, thickset structure as in the Italian remains. The oldest AMH remains found in Romania, Europe looked nothing like Cro Magnons judging by the reconstruction.

The results obtained of a supposed mtDNA CRS in those Italian remains, I have maintained is 1) due to contamination from the numerous modern Europeans who handled the bones from exhumation, storage and study, 2)Due to outright scientific fraud for whatever reasons and motives. Scientific fraud is not an uncommon occurrence, as is unfortunately other antisocial behaviour like theft.

All Europeans of whatever haplogroup originate in peoples who emigrated to Europe well after the advent of farming, and contain only a few indicators of the previous populations whether Mesolithic hunter gatherers or Neolithic farmers.

If it looks like a duck it quacks like a duck and plucks like a duck. It is a duck.

12308A eliminates U.

00073A,11719G

Puts it in HV and subgroups. Almost all the groups in R and HV have a HVR1 mutations. Most of the papers we are looking at have lesser standards for Haplogroup identification.

This obsession about this paper is irrational. Argue with me that it is just one person if you must, that is at least valid. Argueing about H is unseemly. It is H. Almost everyone but you believes this. The alternative is academic fraud. And subclades of H are over 30,000 years old. Hunter -Gatherers.

No U samples. Perhaps by a very long shot Paglicci-23 is U*, but definitely not U4/U5. Put this together with the Paleolithic results from Iberia and Morocco and there's a serious problem with the notion that all of Europe was U4/U5 and became replaced by the H farmers; who actually had more N1a than H, which has somehow magically disappeared, while the U4/U5 of the displaced aborigines is today 50 times more abundant than N1a! You can't say the demic diffusion theory has been conclusively proven with all these flagrant contradictions still existing. I can admit that genetic continuity has its own very serious contradiction due to the U4/U5 mtdna results.

Those are probably including HVRII. A better gauge would be just from HVRI, because all these ancient mtdna studies report only HVRI mutations, so here are the frequencies of CRS as per Richards (2000), using only HVRI mutations (the range is limited to 16090-16365, but I've verified in mitosearch that there's no big difference if the range is extended to include up to 16519):

By the way, no conclusions can be drawn about the Lapps using mtdna because they have unquestionably undergone genetic drift. 50% of their mtdna is U5b1b1 (something like that), and they also have 30% or more V, which is completely off the charts compared to any other population of the world, a situation similar to Ashkenazis having 30%+ mtdna K, while the rest of the Caucasian world ranges between 3% and 10%.

Lapps have 0.0% U4 and 0.7% U5a (Simoni, 2000 [n=240]; Tambets, 2004 [n=445]). If their relative proportions of U4, U5a, and U5b in the pre-drift population were similar to Finns, then a very rough estimate of their pre-drift U5 frequency results in 7% to 10% U5.

Dienekes' assertion that mtdna frequencies have changed noticeably in the last 2000 years (Etruscans, Danish, England) doesn't jive with the fact that today's mtdna frequencies are notoriously homogenous across Europe, which would require either very massive internal migrations in the last 2000 years, to the point of almost complete population replacement everywhere in Europe, or an amazing coincidence (all the random fluctuations just happened to randomly result in each region of Europe having almost the exact same haplogroup distribution). Or a third possibility... evolution... which would have to be on an unprecedented scale, and would involve a bizarre outcome consisting of several dozen haplogroups, each having a specific evolutionary advantage so as to explain why there isn't a single dominant haplogroup in Europe but instead a basket of many. And their evolutionary advantages would have to act upon pan-European variables, not stuff like climate or diet, otherwise Mediterranean, Atlantic, and Continental regions should have resulted in different sets of haplogroups; instead, they're all basically identical, with minor differences.

1. Pitted Ware is a Neolithic culture (albeit regressed to hunter-gathering) of the Chalcolithic Age (paneuropean chronology), including large areas of the Eastern Baltic and central Sweden. It has nothing to do with Denmark.

2. Neolithic did not arrive to Denmark with Pitted Ware or its somehow related (both seem originated somewhat in Dniepr-Don) Funnelbeaker culture. Neolithic in Denmark and Scania is a matter of an older period and culture: Ertebölle, a local development with whatever influences from Danubian Neolithic. Denmark-Scania had been in the Neolithic since long before these Ukranian influences.

3. Ertebölle evolved, apparently on its own, into early Funnelbeaker (TRBK-B) but in later TRBK (A, C and late) it is apparent the same kind of "Ukranian" influences as those of Pitted Ware, all suggestive of a migration from Eastern Europe into the Baltic Area (earlier than Kurgans). At that time Denmark also experimented the western influence of Megalithism, showing a hybrid culture that persists for about a millennium until the Single Burials period (Kurgan, Corded Ware).

I did counter your argument. But you just dont seem to see what you dont want to see.

Your counter-argument amounts to this:

"HV*, well that is a bit broad and unspecified. But most known have HVR1 mutations."

"Most known" is vague. Most H has HVR1 mutations too. And, the percentages that do not, and are simply rCRS were given by me above. And, the reason why rCRS is usually thought to be H today is because H is much more frequent than HV today. But, you can't make that inference about 28,000 years ago.

So your counter-argument is non-existent. In order to infer that Paglicci was haplogroup H, you need to assume that H was more frequent than HV* in Europe 28,000 years ago. So, you prove H's presence in European hunter-gatherers by assuming it. Faulty logic.

For the record Greece and Italy have the lowest frequencies of H in Europe. NOT what you would expect if there was an East to West migration of H into Europe via Greece.

There are much better data for Greece than the ones in the Eupedia article.

Also, there is no need to use modern frequencies of H, when we now have dozens ancient pre-farming samples from Europe, where the frequency is much much lower than today. It is also strange to make inferences about directionality of movement based on a few percentage points of modern differences, when we observe 40%+ of difference between pre-farming and modern Europeans.

I insist: Pitted Ware is not Paleolithic and is contemporary of TRBK with Megalithism, though in different geographical areas.

There's absolutely no reason to think that Pitted Ware peoples, surely just arrived to the area from Ukraine in a rather well known migratory wave, were older in the area than the people more or less reflected in the Megalithic burial sample. In fact the opposite is probably true.

These maps were made by me using 25,000 samples from scientific studies (and in the case of U5 an extra 10,000 samples from mitosearch). It's as detailed as it gets. Both maps are a picture perfect description of genetic continuity. The results are exactly what would be expected from genetic continuity.

U5 gradually declines away from its center, as happens with all haplogroups everywhere in the world. There are no abrupt, sharp changes, except where there are serious geographic barriers, which is in line with ancient genetic continuity (historic movements of people were much less affected by sea and mountains).

According to the latest mtdna results, Europe would have been 70% or more U5, and was then swamped by Anatolian/Mid-Eastern farmers who had very little U5, if any at all. There should be some very sharp clines in the modern map of U5. Instead, U5 shows the typical pattern of all haplogroups everywhere in Africa and Eurasia: it gradually declines from 10% in East Europe to 7% in southeast Europe, to 3% in Anatolia, to 1% in the Middle East. You see the same rate of decline as you travel east to the Urals, then Central Asia, and then eastern Siberia. Or west into West Europe and then into North Africa. And notice that Greece and Bulgaria have pretty much the same frequency of U5 as Iberia or Britain, despite that the farmers entered through the southeast. The Caucasus, much closer to Anatolia, has a rate of U5 around 7% or 8% (n=thousands of samples), comparable to West Europe. That doesn't quite fit with the farmer expansion theory.

By the way, there was an ancient mtdna study of Mycanean Crete (3500 ya) published last year that found 1 U5a1, 1 K (2 samples, brother and sister), and 1 undefined that could have been almost anything, including H and U4, but excluding U5.

Dienekes mentions that U4/U5 results like if the case has been closed, but he conveniently forgets the paleolithic Iberian results (making the ridiculous claim that it's not trustworthy), and he conveniently ignores the huge problem of the Neolithic results, which is their massive presence of N1a. With regards to N1a, he assumes evolution (what a trump card) must have made it disappeared, with no evicence. The same assumption he's accusing Annie Mousse of displaying. When I noted the European Neolithic farmers had already been farming for several thousand years and should have lost their N1a already, Dienekes makes another assumption and says that obviously N1a must have been more abundant at the beginning of the farming revolution and was on its way down in the European Neolithic farmers.

The U4/U5 results are very interesting. They're a huge spoke in the wheels for genetic continuity. But the farmer expansion theory is riddled with its own very serious contradictions, which Dienekes assumes away with evolution and without any evidence. Proportionately, the reduction of N1a was a lot worse than the reduction of U4/U5, to the point that when Haak published his study of the mtdna of Neolithic Europe with 25% N1a, he concluded that modern Europeans weren't descended from Neolithic farmers. I don't agree with his logic, but it shows how the farmer theory has just as many very serious contradictions as the U4/U5 studies have now thrown onto the genetic continuity theory.

These maps were made by me using 25,000 samples from scientific studies (and in the case of U5 an extra 10,000 samples from mitosearch). It's as detailed as it gets. Both maps are a picture perfect description of genetic continuity. The results are exactly what would be expected from genetic continuity.

Right, 25,000 samples from scientific samples, and 10,000 samples from mitosearch, all of them from the late 20th/early 21st century, prove genetic continuity for tens of thousands of years...

With regards to N1a, he assumes evolution (what a trump card) must have made it disappeared, with no evicence.

Do me the courtesy of reading what I write before repeating the same nonsense.

I will say it once more, but, given your close-mindedness and a priori belief in a Paleolithic origin of Europeans, I doubt that it will do any good.

Primary descent from Neolithic farmers can be reconciled with the modern gene pool if:

1. N1a experienced a reduction in frequency.

Primary descent from Paleolithic hunter-gatherers can be reconciled with the modern gene poll if:

1. U- subclades experienced a massive reduction in frequency.2. Non-detected haplogroups in the Paleolithic gene pool were in fact present.3. These multiple not detected haplogroups experienced a massive increase in frequency.

ERGO: Descent from Neolithic farmers is a much simpler hypothesis than descent from Paleolithic hunter-gatherers.

PS: It doesn't matter one iota if you include the questionable late Portuguese Mesolithic population, as the total sample remains U-dominated and H-deficient.

PPS: The hypothesis of neutral evolution, i.e., the derivation of the modern gene pool from that of the hunter-gatherers via simple genetic drift has been tested and rejected. So, if you accuse me of postulating "natural selection", you should be aware that natural selection is the only way in which the Paleolithic gene pool could have transformed itself into the modern European gene pool.

I'm showing the massive evidence in favor of genetic continuity, not to prove it, but to demonstrate that your idea that these studies have settled the matter and constitute a nail-in-the-coffin of the genetic continuity theory are way overblown. [you forgot the 3 Paglicci samples, no U4/U5, but 2 of them are in superhaplogroup HV]

The most interesting part of these papers is that they show the dominance of U in Northern Europe AFTER the glaciation period (and before). This implies that either Humans remained (with much diminished numbers) in Northern Europe during the last ice age, or repopulated Northern Europe from a different ice age refuge. Fascinating either way.

Definitely. Almost all of the H and U haplogroups are so old that finer details of the subclades are the only way to make progress.

Also, there seem to be an enormous number of founder effects since the original settling of Europe and continuing through until our most recent past. With so many different haplogroups and subclades "competing", and likely thousand of local founder effects throughout the millennia, I am not surprised that many strains did not or almost not survive - some of their numbers in absolute terms - especially before the advent of agriculture - must have been minute, but with a surprisingly high initial diversity.

That they were (regressed into) hunter-gatherers doesn't mean necessarily that they represent local Paleolithic continuity. This also applies to the other paper, where most of the samples (all the ones from Poland and Latvia) also belong to this "regressive" Neolithic cultural area.

If you have to make a case for pre-Neolithic peoples of Northern Europe being different, you'd have to first of all sample actual pre-Neolithic peoples that offer some guarantees. To me this only seems to suggest that the U5/U4 lineages may have arrived from Neolithic Ukraine within this migration. If confirmed, it'd mean that Dniepr-Don and Danubian Neolithics were very different genetically.

Here there is an interesting article on the Scandinavian facies of Pitted Ware: they had pigs and occasionally farmed cereals!

The rounded-pointed base of the pots, among other items (burial customs) that they came from Dniepr-Don offshots, that at the time were colonizing the NW (Belarus, Baltic countries...) similarly in a regressive way to hunter-gathering, a practice that was never fully abandoned in Neolithic Ukraine.

I've been paying attention. I have to say, my knowledge of prehistoric cultures is almost zero, terms like Gravettian dumbfound me completely, I couldn't place them anywhere on a map or a timeline, I don't have the slightest clue. Some questions...

I'm surprised at the possibility that a population could regress back to hunter-gatherer. Do you have an explanation how/why that would happen? [Climate gets worse? That's all I can think of.]

We would still have the issue of these populations having vastly huge frequencies of U4/U5. The contradictions with the Iberian/Paglicci results, or with the amazingly homogenous modern mtdna distribution, would still stand.

So when did the first farming (aka Neolithic?) really occur in Denmark/Sweden?

There were a number of cultures in the Neolithic. Near as I can see if they are rich in Haplogroup U they are considered Hunter-Gatherers and if there are a lot of other haplogroups they are considered farmers.

So the Pitted Ware people (who farmed) are considered regressed farmers as they are rich in Haplogroup U.

And the Funnel Beaker people are considered farmers in the Sweden paper (contains no U) and Hunter-Gatherers in the Bramanti paper (rich in U).

Techniques like these ensure that the paper comes to the desired conclusions (that is, farmers physically replaced Hunter gatherers). In the teeth of evidence to the contrary.

The subtext is the implication that the Hunter-Gatherer Haplogroups were genetically inferior. That they were replaced by a more advanced model of people with the cultural change (farming).

Personally I think reality is rarely that simple. Betamax and VHS comes to mind for example. And in the hunting/farming transition there is plenty of evidence of U embracing farming cultures.

As to the hunter-gatherers' "regression", I would say that the farther back you go in time, the more professions become parallel cultures.

Nowadays, most people have a long general education followed by a short specialized one, followed by stints at one to a few different professions. And most people use similar tools and live similar lives.

Going back just a few hundred years, apprentices traveled hundred of miles, lived with their masters, and learned "secrets" of trade, tools and construction that were not easily obtainable otherwise.

Then you had other "business models" like gypsies and traveling salesmen. Going back farther, perhaps brewers or tradespeople had rather different cultures than their hosts (one interpretation of the Bell-Beaker culture).

Also, note that these "hunter-gatherers" were predominantly fishers. Fishermen and trappers have had very successful "business models" while living quite different lives, for millenia and up until the most recent past (even in the US, and including e.g. the Inuits).

The fishing people could have also had uncharacteristically dark skin and dark hair for Nordic latitudes, making genetic separation easier (because they looked so different).

I'm surprised at the possibility that a population could regress back to hunter-gatherer. Do you have an explanation how/why that would happen? -

For what I know, it's mostly that people never really stopped hunting and gathering, that agriculture and hunter-gathering were complementary in Neolithic Ukraine/Don, though with a tendency to increase the agricultural/herding fraction of the economy.

Then the NW area (relative to Ukraine: Baltic) was also one of low population densities (excepting maybe Denmark and some specific LBK districts): it was a "frontier" area where lot of wild nature stil existed making hunter-gathering more efficient than an agriculture original from the Mediterranean that always had problems adapting to high latitudes, specially the cold/wet Oceanic climate.

I guess it's about that. You could well say that fur trappers in other contexts were hunter-gatherers, even if they belonged to larger neolithic economic units/societies. Probably fur trapping and maybe even trading these was a big deal of their economy, in a time when trade was already going on at large scale (that's why it's Chalcolithic and not mere Neolithic, even where soft metals were not used locally).

Ask an expert anyhow.

We would still have the issue of these populations having vastly huge frequencies of U4/U5. The contradictions with the Iberian/Paglicci results, or with the amazingly homogenous modern mtdna distribution, would still stand.

Indeed. But it makes some sense because U (U5 and even more others like U4, U3) tends to be more frequent towards the East, while H is instead towards the West.

So when did the first farming (aka Neolithic?) really occur in Denmark/Sweden?.

According to my notes, in the second half of the 5th milennium, with Ertebölle culture, which was largely oriented to sea exploitation but had already clear agriculture, cattle and pottery. By the 4th milennium it evolved into an early Funnelbeaker form (TRBK-B, which is older than TRBK-A - there was an error in early estratigraphy, it seems). And then, in the advanced 4th milennium is when the Dniepr-Don related hunter-gatherers of Sarnowo and what some also call Pitted Ware of the East Baltic begin migrating, which is roughly coincident with Seredny-Stog II, that is a mixed DD/Kurgan complex rather than a single culture. The Eastern Pitted Ware phenomenon would have influence, for what I've read the Danish-plus TRBK in the later phases, while they were also influenced by the west by Megalithic culture: this caused a "double" cultural area, with two types of burials: megalithic ("collective" or "clannic") and "standard" (with strong resemblances to DD: ochre and stuff like that). In middle coastal Sweden Pitted Ware strictu sensu dominates instead almost solo.

But read other stuff because it's not like I'm uber-informed on this issue anyhow.

As the few last comments point it, I don't see much of archeological-related arguments in all these ideas about Hunter-Gatherers & Farmers... I see people with an idea, trying to say that whatever genetic data implies whatever occupation. It is not really an anthropology blog. It is an anthropology blog focused on genetic discoveries. Yet very interesting.

This comment is a little late and slightly off topic, but I can't help but notice how often cranial shape is used to bolster genetic relatedness arguments on your blog.

The shape of the human head is affected primarily by two non-genetic forces--the toughness of the foods people eat while their jaws are developing and the actual physical molding/shaping of thier heads done by the doctor or midwife immediately after their birth (all babies pop out with their heads a little misshapened from their trip down the birth canal--studies have shown that cultural preferences in baby-head-shape influence how baby heads are molded post-birth--in the US, baby-head-shape has change from a rounder one to a more oblong one, just since 1940s/50s--and this is 100% the result of post-birth molding done at the hospital).

Proper formation of the human jaw requires chewing on fairly tough substances for the first 18-20 years of life. The result is a broad/wide face. The long, narrow, Anglo-European face is actually not genetic at all but the result of eating soft food as children (and actually fairly typical of all soft food cultures)--it is and incorrect jaw formation and a mistake that results in widespread malocclusion (most people think malocclusion--i.e. crooked teeth-- is genetic, but it rarely is, it is predominantely dietary).

The fact that this facial type emerged with the onset of agriculture was not a coincidence. It is cause and effect. The human diet became considerably softer after the advent of agriculture and thus the narrow, badly malocclused face was "born."

To correct for the soft diet today, you have to use a crozat retainer--even adults with fully matured jaws can benefit from a crozat retainer. In comparisons of photographs of twins, where one twin was given traditional braces and teeth removal to correct for malocclusion, and the other a crozat retainer, most people found the wider, crozat face much more attractive, and determinations of attractiveness is a proxy for human assumptions of genetic fitness.

This comment is a little late and slightly off topic, but I can't help but notice how often cranial shape is used to bolster genetic relatedness arguments on your blog.

Human craniometric traits are heritable, so they _can_ be used to bolster genetic relatedness arguments, provided that a sufficient number of them are used, in which case the clustering produced using autosomal markers is largely reproduced.

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